FUNCTION OF PLAY IN BEHAVIORAL ENRICHMENT OF CAPTIVE ASIAN ELEPHANTS (ELEPHAS MAXIMUS) By CONNIE K. DUER Bachelor of Arts in Natural Science Avila University Kansas City, Missouri 1989 Master of Science in Biology Missouri State University Springfield, Missouri City 2004 Submitted to the Faculty of the Graduate College of the Oklahoma State University in partial fulfillment of the requirements for the Degree of DOCTOR OF PHILOSOPHY July, 2009 FUNCTION OF PLAY IN BEHAVIORAL ENRICHMENT OF CAPTIVE ASIAN ELEPHANTS (ELEPHAS MAXIMUS) Dissertation Approved: Dr. Charles I. Abramson Dissertation Adviser Dr. Larry G. Talent Dr. Anthony A. Echelle Dr. Timothy J. Bowser Dr. A Gordon Emslie Dean of the Graduate College ii ACKNOWEDGMENTS “Knowledge and understanding are life’s faithful companions who will never prove untrue to you. For knowledge is crown, and understanding your staff; and when they are with you, you can posses no greater treasure.” Kahlil Gibron First and foremost I want to thank God for the power of Prayer and for all those dear souls who practice it. Without them this manuscript would not have been possible. May I extend my heartfelt appreciation to the succeeding organizations and individuals? Each has played an essential role in making this research possible. My thanks to the Dickerson Park Zoo and the Friends of the Zoo and their management and staff for the opportunity to carry out our research. Particular recognition and thanks to Melissa Carden, Ph.D. and Nicole Conover whose diligence made this study possible. I extend my gratitude to Oklahoma State University and all the many people who work rigorously to help each student overcome obstacles and achieve their goal. My sincere appreciation to the Zoology Department as well as the Graduate College for their support and guidance. I would be remiss if I did not give special recognition to Dr Talent, Dr. Echelle, and Dr. Boswer for their expertise and willingness to hang in there with me when it would have been easier not to. If there were such as thing as The Metal of Honor for valor and perseverance above and beyond the call of duty for a major advisor it would have to be given to Dr Abramson. There are not enough words to express my appreciation for his dedication to iii getting me through. He started the journey with me and honored his word to see me through. An abundance of appreciation to my friend and Son-in-Law, Dave Luedde, for his friendship and encouragement. He made the difference so many times. To my daughter Stephani Luedde who has believed and hoped when I couldn’t; I send my love and gratitude. Last but far from least, I acknowledge the elephants who allow us to observe and record their lives. I can only hope that in some small way we have made their lives better. Dr. Rasmussen, know that your light has not gone out. iv Dedicated to Onyx “Thus spoke the speechless animal whom my heart had understood.” Kahlil Gibran v TABLE OF CONTENTS Chapter Page I. A COMPREHENSIVE PREFACE TO LIVING MEMBERS OF ORDER PROBOSCIDAE ............................................................................ 1 Chapter Rationale ...................................................................................1 Fossil Record and Evolution of the elephant……………………. ........1 Superorder Paenungulata .....................................................................6 Superorder Tethytheria (“beasts of the Tetys”) .....................................7 Family Elphantidae: Loxodonta and Elephas ......................................11 Present Day Elephant Population in Asia ............................................14 Reproductive and Social Behavior of Asian Elephants in the Wild ....16 Summary ..............................................................................................20 II. ACCOUNT OF ANIMALS IN CAPTIVITY AND MANAGEMENT OF CAPTIVE ASIAN ELEPHANTS (ELEPHAS MAXIMUS) ................. 22 Chapter Rationale ............................................................................... 22 History of Zoos .................................................................................. 22 Modern Day Zoos .............................................................................. 27 Differentiation of “Animal Rights” and Animal Welfare” ................ 28 History and Formation of the American Association of Zoological Parks and Aquariums (AZA) ............................................................. 30 North American Asian Elephant Population ...................................... 32 Husbandry of Asian Elephants in North American Zoos .................. 33 Meeting the Challenges of Captive Breeding .................................... 36 Summary ............................................................................................ 38 III. AFFILIATION OF STRESS AND WELL-BEING IN CAPTIVE ANIMALS .................................................................................................. 39 Chapter Rationale ............................................................................... 39 Stress Defined .................................................................................... 39 Nature of Stress .................................................................................. 40 Acute and Chronic Stress Response and its Outcome ....................... 41 Coping Mechanisms and Stereotypic Behavior ................................. 43 Adaptive Features of Stress ............................................................... 46 Comparing the Captive Animal to its Wild Counterpart ................... 48 vi Chapter Page Animal Welfare or Well-being .......................................................... 50 Coping: A Gateway to Well-being .................................................... 52 Assessment of Well-being ................................................................. 53 Summary ............................................................................................ 55 IV. EFFECT OF ENRICHMENT IN NORTH AMERICA ............................. 56 Chapter Rationale ............................................................................... 56 Enrichment Defined ........................................................................... 56 Environmental and Behavioral Enrichment Defined ......................... 56 Purpose and Implementation of Enrichment ...................................... 57 Enrichment Timeline ......................................................................... 58 Approaches to Environmental Enrichment ........................................ 62 American Association of Zoological Parks and Aquariums (AZA) Addresses Enrichment Concerns ....................................................... 65 Enrichment Put into Practice ............................................................. 66 Summary ............................................................................................ 67 V. ASSESSMENT OF ANIMAL PLAY AND POTENTIAL UTILIZATION OF PLAY AS BEHAVIORAL ENRICHMENT FOR CAPTIVE ASIAN ELEPHANTS (ELEPHAS MAXIMUS) IN NORTH AMERICAN ZOOS .................................................................................... 68 Chapter Rationale ............................................................................... 68 Play Defined....................................................................................... 68 Learning: A Component of Play ........................................................ 70 Object Play ......................................................................................... 71 Locomotor Play .................................................................................. 72 Social Play ......................................................................................... 74 Cost of Play ........................................................................................ 75 Prologue to Research Inquiry ............................................................. 76 VI. METHODS ................................................................................................. 78 Chapter Rationale ............................................................................... 78 Study Subject, Asian Elephant, (Elephas maximus) .......................... 78 Animal Housing ................................................................................. 79 Diet ..................................................................................................... 79 Behavioral Data ................................................................................. 79 Description of Enrichment Items ....................................................... 80 Blood Collection ................................................................................ 81 Statistical Analysis ............................................................................. 81 Calculations........................................................................................ 82 vii Chapter Page Description of Elephants and Play Groups ........................................ 83 Summary ............................................................................................ 85 VII. RESULTS ................................................................................................... 86 Chapter Rationale ............................................................................... 86 Age ..................................................................................................... 86 Vocalizations.....................................................................................